The Condor88:478A82 0 The CooperOrnithologicalSociety1986

A FOSSIL MOTMOT (AVES: MOMOTIDAE) FROM THE LATE MIOCENE OF FLORIDA’ JONATHANJ. BECKER Division of Birds, National Museum of Natural History, SmithsonianInstitution, Washington,DC 20560 Abstract. A fossil motmot, known from the proximal end of a humerus, from the late Miocene (early Hemphillian, approximately 8 MYA) Haile XIXA locality, Alachua County, Florida, is morphologicallyindistinguishablefrom that of living speciesof Electron, Momotus, and Eumomota. This is the first fossil record of the family Momotidae in the Tertiary of the New World. The presenceof a motmot in Florida prior to the land connection between North and South America provides evidence for the Momotidae being widely distributed in the southern part of North America during the late Tertiary, then invading South America during the “Great American Interchange.” Key words: Aves; Coraciiformes;Momotidae; Florida; Miocene; Haile XIXA; zoogeography; Great American Interchange.

INTRODUCTION The Momotidae is a distinctive family of Coraciiformes whose modem distribution is restricted to Central and South America. The fossil record of this family is meager, as until now the only Tertiary record of a motmot was Protornisglurniensisfrom the Oligocene of Switzerland (Olson 1976). Living speciesare known from a few prehistoric sitesin the Yucatan Peninsula and Brazil (Brodkorb 197 1). MATERIAL AND METHODS Humeri of each speciesof momotid were examined: Hylomanesmomotula,Aspathagu-

lark, Electronplatyrhynchum, E. carinatum, Eumomotasuperciliosa, Baryphthengus ruficapillus,B. martii, Momotusmexicanus,and M. momota.Representative humeri of all other families of Coraciiformes and other “higher non-passerines”were also examined. Anatomical terminology follows Baumel et al. (1979). Measurements, defined in Table 1, were taken using dial calipers accurate to the nearest0.05 mm and then rounded to the nearest 0.1 mm. Abbreviations: MYA = million years ago, UF = Florida State Museum, UK = University of Kansas, UM = University of Michigan, USNM = National Museum of Natural History, Smithsonian Institution. SYSTEMATIC

PALEONTOLOGY

ORDERCORACIIFORMES: FAMILY MOMOTIDAE The proximal end of the humerus of Momotidae may be distinguishedfrom all other Coraciiformes in having the following combination of characters: 1) attachment of the dorsal head of the M. humerotriceps expanded dor-

I Received 8 January 1986. Final acceptance 18 February 1986.

sally, causingthe margo caudalis (i.e., capitalshaftridge of Bock 1962) to be directed toward the tuberculum dorsale and 2) attachment of the M. pectoralisprimarily restrictedto a small triangular area on the crista pectoralis (seeFig. 1). MOMOTIDAE,GENUSAND SPECIES INDETERMINATE

Material. Proximal end of left humerus, UF 6 1400, Vertebrate Paleontology collection of the Florida State Museum. Collected on 21 January 1984 by Mr. Phillip M. Whisler of Venice, Florida (original number PMW 15 17) and donated to the Florida State Museum. The specimen is light brown in color and is permineralized. The preservationis similar to that of other early Hemphillian taxa from this locality. Locality.The Haile XIXA locality was exposedin a limestone quarry 4 km northeast of Newberry, Alachua County, Florida (NE l/4, Sec. 26, T. 9 S., R. 17 E., Newberry Quadrangle, U.S. Geologic Survey 7.5 min series topographical map, 1968). Sediments are fluvial and include both “gar-scale” and phosphatic gravels. Fossil mammals from the same stratigraphic level as the above specimen, include Pliometanastes, Osteoborus cf. validus,Geolocidae, Pediomeryxhamiltoni, at least five speciesof Equidae, cf. Hypolagus,GomphoAphelops,Procamelus, theriidae, Teleoceras, and Aepycamelus (G. S. Morgan, pers. comm. 1985). The co-occurrenceof thesemammalian taxa, and that of the crocodylian Gavialosuthus, indicate an early Hemphillian North American Land Mammal Age, approximately 8 MYA. Other avian taxa known from this locality include an undetermined species of cormorant Phalacrocorax sp.,an extinct species of anhinga Anhingugrandis,and a few specimens of indeterminate anatids. Additional in-

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TABLE 1. Measurements of the proximal end of the humeri of speciesof Motmotidae. Data are mean, standard deviation, and observed range. Measurements are W-SHAFT, transverse width of midshaft; D-SHAFT, depth of midshaft; W-PROX, transversewidth of proximal end from the external tuberosity (Tuberculumdorsale)to the most ventral face of the bicipital crest (Crista bicipitalis);D-PROX, depth of proximal end, from the bicipital surface(F&es bicipitalis)to the internal tuberosity (Tuberculum ventrale), measured at right angles to the long axis of the shaft; D-HEAD, depth of head, measured parallel to the axis of the head; L-DELTOID, length of deltoid crest (Crista pectoralis),measuredfrom the external tuberosityto the most distal extensionof the deltoid crest.Number of specimens equals 1 for all exceptBaryphthengusruficapillus(n = 7), M. mexicanus(n = 3), and M. momota (n = 15). B. rujicapillus includes specimensof both B. rujicapillusmartii and B. rufcapillus semirufi. Species

Momotid, UF 6 1400 Hylomanes momotula Aspathagularis Electronplatyrhynchum E. carinatum E. superciliosa Baryphthengusrujicapillus

Momotus mexicanus

M. momota

W-SHAFT

D-SHAFT

2.8 2.1 2.4; 2.6 2.9; 2.9 2.8 2.7; 2.9 4.23 0.21 4.0-4.6 3.34 0.38 3.0-3.4 3.60 0.32 3.2-4.2

2.5 1.8 1.9; 2.2 2.4; 2.6 2.3 2.6; 2.4 3.80 0.20 3.5-4.1 2.80 0.23 2.6-3.0 3.17 0.21 2.8-3.6

formation on this and other coetaneouslocalities in Florida is found in Becker (in press). Description. The humeri of the living species of Momota, Eumomota,and Electronare inseparable from one another by size or qualitative characters.The fossil specimen is morphologicallyidentical with motmots ofthis size. It may be distinguishedfrom the humerus of speciesof Aspathaand Hylomanesin having the attachment for the dorsal head of the M. humerotriceps less expanded dorsally and in having a deeper sulcus lig. transversus, and in being lesspneufrom that of Baryphthengus matic. Additionally, the fossil specimenis easily distinguishedfrom the last three genera on the basis of size (Aspathaand Hylomanes smaller, Baryphthengus larger; Table 1). DISCUSSION Olson (1976a) noted the Old World origin of all Coraciiformes, with the possibleexception of the Todidae. Recent work by Mourer-Chauvire (1982), documenting the presenceof Todidae in the Eo-Oligocene Phosphorites du Quercy, France, confirm that the New World distribution of both the Momotidae and the Todidae is relictual. When and how the families Momotidae and Todidae crossedfrom one continent to the other is not known, but dispersal possibly occurred during the Eocene acrossthe North Atlantic land connection between Europe to North America (DeGeer route). At this time over one-half of the known genera of fossil mammals are shared between North America

W-PROX

9.1 6.8 7.7; 8.2 9.0; 9.2 9.1 8.8; 9.2 13.04 0.57 12.6-13.5 10.05 0.40 9.7-10.5 11.06 0.74 9.9-12.6

D-PROX

D-HEAD

L-DELTOID

3.6 2.5 3.3; 3.5 3.5; 3.6 3.8 3.5; 3.7 5.61 0.35 5.3-6.3 4.13 0.35 3.9-4.5 4.86 0.35 4.4-5.5

2.7 2.0 2.3; 2.4 2.8; 2.9 2.7 2.7; 2.7 4.26 0.24 4.0-4.7 3.00 0.24 2.8-3.3 3.60 0.25 3.3-4.1

8.7 6.2 7.9; 8.2 8.9; 9.8 9.0 9.2; 9.1 13.19 0.50 12.5-13.9 10.19 0.55 9.8-10.8 11.48 0.90 10.1-13.2

and Europe (McKenna 1975). The climate acrossthis area was warm temperate as shown by the paleobotanical record and the presence of amphibians and reptiles (West et al. 1977, West and Dawson 1978). During the Mid-Tertiary, the Gulf Coast and easternNorth America retained its moist subtropical vegetation(Graham 1964, Webb 1977) and remained relatively unaffected by the development of a midcontinental savannaprovince that extendedwell southonto the Mexican Plateau. By the early Miocene, a “Gulf Coast Chronofauna” (Tedford et al. in press)developed in what is now the southernpart of North America, reaching from Florida (Webb 198 l), west acrossto Texas (Wilson 1956) and south to Panama (Whitmore and Stewart 1965). The middle Miocene saw the breakdown of this “Gulf Coast Chronofauna” and the establishment of a fauna1continuity between localities in Texas, Florida, and the Great Plains. A savanna corridor had expanded eastward, reducingthe subtropicalforest and finally breaking the connection between the mesic forests of the southeasternUnited Statesand those of the tropics (Webb 1977). The breakup of the mesic forestsof the Gulf Coast is reflected in the distribution of living and fossiltaxa. Several vertebrate groupspresently restricted to the Neotropics are representedin the fossilrecord acrossthe Gulf Coast in the late Miocene (Estes 1970, Savage 1966, Olson 1976b, Becker, in press). The joining of the North and South American continents,and the biotic interchangethat

FOSSIL MOTMOT FROM FLORIDA

took place between them is well-documented (Stehli and Webb 1985) generally under the rubric “Great American Interchange.” Mammals provide the most informative fossil evidence for the timing of these events. Fossil mammals show two distinct waves of interchange in the Neogene between North and South America. A very limited overwater exchange occurred in the late Miocene (early Hemphillian) that included South American megalonychid and mylodontid sloths and a North American procyonid (Marshall et al. 1979). At this time, the distance between the two continents is estimated to have been 150 km or more (Webb 1978). Later, about three million years ago,the land connection between North and South America wascompleted, permitting a much greater exchange (Stehli and Webb 1985). Motmots are nonmigratory, sedentarybirds. They have a limited ability to cross large expansesof open water, as shown by their absencein both the fossil and recent record from all Caribbean islandsexcept the two (Trinidad and Tobago) that were connected to South America during the Pleistocene.It is therefore unlikely they would have crossed the water barrier that existed between North and South America in the late Miocene. The presenceof a motmot in Florida during the late Miocene, the probable inability of this family to cross the water gap that existed at this time between North and South America, and the probable homogeneity of habitat between Mexico and Florida, all argue that the motmots were widely distributed across the southern part of North America at that time. It wasprobably not until a completeland bridge formed between the continentsthat momotids entered South America, much as Chapman (1923) proposed, based on his study of the distribution of living speciesand subspeciesof the genusMomotus. The family Momotidae probably diversified in the southern part of the North American continent, and then invaded South America after the closure of the Panamanian Seaway about three million years ago, providing an avian example of the “Great American Interchange.” ACKNOWLEDGMENTS For the loan of modem and/or fossil specimens,or access to collections,I thank S. D. Webb, B. J. MacFadden, T. Webber, J. W. Hardy, Florida StateMuseum; P. Brodkorb, University of Florida; R. and M. Mengel, University of Kansas;R. B. Payne, University of Michigan; S. L. Olson, R. L. Zusi, National Museum of Natural History, SmithsonianInstitution. This manuscript was greatly improved by discussionswith S. L. Olson and G. S. Morgan. I am

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grateful to S. L. Olson, R. L. Zusi, G. S. Morgan, D. W. Steadman, and S. D. Webb for their comments on this manuscript. Phil Whisler of Venice, Florida very generouslydonatedthe specimenof fossilmotmot to the Florida State Museum. Photographsare by Victor E. Krantz.

LITERATURE

CITED

BAUMEL,J. J., A. S. KING, A. M. LUCAS,J. E. BREAZILE, AND H. E. EVANS. 1979. Nomina anatomica avium. Academic Press,London. BECKER,J. J. In press. The fossil birds from the late Miocene and early Pliocene of Florida. I. Geology, correlation, and systematicoverview. Centre Regional de Publication de Lyon. BOCK,W. J. 1962. The pneumatic fossaof the humerus in the Passeres.Auk 791425-443. BRODKORB, P. 1971. Catalogue of fossil birds, Part 4 (Columbiformes through Piciformes). Bull. FL. State Mus. Biol. Sci. 15:163-266. CHAPMAN,F. M. 1923. The distribution of motmots of the genus Momotus. Bull. Am. Mus. Nat. Hist. 48: 27-59. ESTES,R. 1970. Origin of the Recent North American lower vertebrate fauna: an inquiry into the fossil record. Forma Functio. 3: 139-163. GRAHAM,A. 1964. Origin and evolution of the biota of southeasternNorth America: evidence from the fossil plant record. Evolution 18:571-585. MARSHALL,L. G., R. F. BUTLER,R. E. DRAKE, G. H. CURTIS,AND R. H. TEDFORD. 1979. Calibration of the Great American Interchange. Science 204:272279. MCKENNA,M. C. 1975. Fossil mammals and the early EoceneNorth Atlantic land continuity. Ann. MO. Bot. Gard. 62:335-353. MOURER-CHAUVIRB, C. 1982._ Les oiseaux fossiles des Phosphoritesdu Quercy (Eocene suptrieur a Oligocene superieur):implications paltobiogeographiques. Geobios (Lyon), memoire special 6:4 13-426. OLSON,S. L. 1976a. Oligocene fossils bearing on the origins of the Todidae and the Momotidae (Aves: Coraciiformes).Smithson.Contrib. Paleobiol.27: 11l119. OLSON, S. L. 1976b. A jacana from the Pliocene of Florida (Aves: Jacanidae).Proc. Biol. Sot. Wash. 89:259264. SAVAGE, J. M. 1966. The origins and history of the Central American herptofauna. Copeia 1966:719-766. STEHLI,F. G., AND S. D. WEBB[EDS.]. 1985. The great American biotic interchange.Plenum PublishingCo., New York. TEDFORD,R. H., T. GALUSHA,M. F. SKINNER, B. E. TAYU)R, R. W. FIELDS, J. R. MACDONALD, J. RENSBERGER,S. D. WEBB, AND D. P. WHISTLER. In press. Fauna1 succession and biochronology of the Arikareean through Hemphillian interval (late Oligocene through late Miocene Epochs), North America. Univ. of California Press, Berkeley. WEBB, S. D. 1977. A history of savanna vertebrates in the New World. Part 1: North America. Annu. Rev. Ecol. Syst. 8:355-380. WEBB, S. D. 1978. A history of savanna vertebrates in

the New World. Part II: South America and the great interchange.Annu. Rev. Ecol. Syst. 9~393-426. WEBB,S. D. 1981. The Thomas Farm fossil site. The Plaster Jacket 37:6-25. WEST,R. M., M. R. DAWSON, ANDJ. H. HUTCHISON.1977. Fossilsfrom the PaleogeneEureka Sound Formation, N.W.T., Canada: occurrence,climate and paleogeographic implications. In R. M. West [ed.], Paleontology and plate techtonics,Milw. Public Mus. Spec. Publ. in Biol. and Geol. 2~77-93.

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WEST, R. M., AND M. R. DAWSON. 1978. Vertebrate paleontologyand the Cenozoic history of the North Atlantic region. Polarforschung48: 103-l 19. WHITMORE,F. C., JR., ANDR. H. STEWART.1965. Miocene mammals and Central American seaways.Science 148:180-185.

WILSON,J. A. 1956. Miocene Formations and vertebrate biostratigraphicunits, Texas Coastal Plain. Am. Assot. Pet. Geol. Mem. 40:2233-2246.

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